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Knoetschkesuchus Langenbergensis Gen. Nov. Sp RESEARCH ARTICLE Knoetschkesuchus langenbergensis gen. nov. sp. nov., a new atoposaurid crocodyliform from the Upper Jurassic Langenberg Quarry (Lower Saxony, northwestern Germany), and its relationships to Theriosuchus Daniela Schwarz1☯*, Maik Raddatz1¤a, Oliver Wings2☯¤b a1111111111 1 Museum fuer Naturkunde Berlin, Leibniz Institute for Evolutionary and Biodiversity Research, Berlin, a1111111111 Germany, 2 Department of Natural Sciences, Landesmuseum Hannover, Hannover, Germany a1111111111 a1111111111 ☯ These authors contributed equally to this work. a1111111111 ¤a Current address: Independent Researcher, Halle, Germany ¤b Current address: Stiftung Schloss Friedenstein Gotha, Museum der Natur, Schloss Friedenstein, Gotha, Germany * [email protected] OPEN ACCESS Abstract Citation: Schwarz D, Raddatz M, Wings O (2017) Knoetschkesuchus langenbergensis gen. nov. sp. We report a new, small-sized atoposaurid crocodyliform from the Upper Jurassic of Langen- nov., a new atoposaurid crocodyliform from the berg, Northeastern Germany. Atoposaurids are small-sized Mesozoic crocodyliforms of Upper Jurassic Langenberg Quarry (Lower Saxony, northwestern Germany), and its relationships to mainly European distribution, which are considered to be phylogenetically close to the origin Theriosuchus. PLoS ONE 12(2): e0160617. of Eusuchia. Knoetschkesuchus langenbergensis gen. nov. sp. nov. is represented by two doi:10.1371/journal.pone.0160617 well-preserved skulls and additional cranial and postcranial remains representing different Editor: Leon Claessens, College of the Holy Cross, ontogenetic stages. 3D reconstructions of a juvenile skull based on micro-computed tomog- UNITED STATES raphy allow the most detailed description of cranial remains of any atoposaurid hitherto pre- Received: February 25, 2016 sented. Our new analysis contradicts previous preliminary assignment of the Langenberg Accepted: September 26, 2016 atoposaurids to Theriosuchus. Knoetschkesuchus gen. nov. is characterized in particular by the presence of two dental morphotypes in the maxilla and dentary, slit-like secondary choa- Published: February 15, 2017 nae within a narrow groove on the surface of the pterygoid, absence of lacrimonasal contact, Copyright: © 2017 Schwarz et al. This is an open presence of an antorbital foramen and an external mandibular fenestra, and proportional access article distributed under the terms of the Creative Commons Attribution License, which characters of the interorbital and intertemporal region. A similar combination of characters permits unrestricted use, distribution, and allows attribution of Theriosuchus guimarotae to Knoetschkesuchus, forming the new com- reproduction in any medium, provided the original bination Knoetschkesuchus guimarotae. Our analysis provides an osteological basis for the author and source are credited. separation of Theriosuchus and Knoetschkesuchus and helps further delineate generic dif- Data availability statement: Data have been ferences in other closely related crocodylomorphs. Our phylogenetic analysis corroborates deposited to Figshare (https://dx.doi.org/10.6084/ m9.figshare.3502811.v2). inclusion of Knoetschkesuchus into Atoposauridae and supports a position of Atoposauri- dae within Eusuchia. Funding: The Volkswagen Foundation generously funded the Europasaurus-Project (grant no. 85 882) and OW in the grant initiative ªResearch in Museums." Competing interests: The authors have declared that no competing interests exist. PLOS ONE | DOI:10.1371/journal.pone.0160617 February 15, 2017 1 / 47 New atoposaurid crocodyliform from the Upper Jurassic of northwestern Germany Introduction The discovery of the dwarfed sauropod dinosaur Europasaurus holgeri made the Langenberg Quarry a fossil locality of world-wide importance [1]. The marly limestones at the Langenberg near Goslar, deposited in a shallow marine inlet or a small marginal basin of the German Late Jurassic Basin [2, 3], yield a high diversity of invertebrate, micro- and macrovertebrate as well as plant fossils [4, 5]. Among these fossils are the remains of marine and terrestrial atoposaurid crocodylians [6±8], which here are described for the first time in detail for the first time. Previ- ous preliminary description placed the specimens in the genus Theriosuchus, but provided only limited evidence for that determination [8]. Atoposauridae is a family of small sized crocodyliforms typically smaller than one meter in body length that was first described by the French palaeontologist Gervais in 1871 [9]. They were included into Crocodylia by the German palaeontologist Zittel in 1890 [10]. Atoposaurid crocodyliforms have a short snout with paired external nares, in relation to the overall skull proportions large orbits and relatively small supratemporal fenestrae in relation to the overall skull proportions, enlarged anterior maxillary teeth, an often reduced or closed antorbital fenestra, slender limbs, and occasionally reduced osteodermal armor [11±17]. According to Buscalioni and Sanz [17], Atoposauridae comprises the five ªcoreº genera Theriosuchus, Alliga- torium, Alligatorellus, Montsecosuchus and Atoposaurus [17, 18]. The monophyly of this group has been questioned [16], because both the skull morphology and the small body size are also a strong ontogenetic signal in morphology. Also, the majority of skeletons of atoposaurids have been found in limestone slabs and were preserved more or less flattened, further complicating morphological interpretation. In phylogenetic analyses, usually only the genus Theriosuchus is coded, sometimes also Alligatorium [19±25]. Theriosuchus itself is probably the best-known and best-preserved taxon within Atoposauridae [26, 27], although a good diagnosis of this taxon is still pending. Several characters separateTheriosuchus from other atoposaurids, which make an assignment to Atoposauridae uncertain [8, 16, 17, 19, 24, 28]. Detailed examination of new finds and thorough descriptions will help to shed new light on the taxonomic status of Theriosuchus. Our re-examination of the atoposaurid specimens from Langenberg assigned to Theriosu- chus pusillus (i.e., DFMMh/FV 200 and DFMMh/FV 605) [8] raised doubts regarding the validity of this determination. The exceptionally well preserved material warranted a more thorough description and a detailed comparison with other species of Theriosuchus. Although the Langenberg remains have been preparated meticulously, parts of the skulls remained in the sediment. Recently, micro-computed tomography (μCT) made it possible to study one of these skulls (DFMMh/FV 605) in more detail. We were able to add a lot of information to the anat- omy of the specimens with the help of three-dimensional (3D) images obtained from the μCT studies. In summary, the ªTheriosuchusº material from Langenberg provides new information on the characters included in a diagnosis of Theriosuchus, adds new information on different ontogenetic states and 3D studies of its anatomy, and in the end makes it possible to separate these specimens from Theriosuchus. Anatomical abbrevations used in figures An, angular; aof, antorbital fenestra; boc, basioccipital; bsph, basisphenoid; bsphr, basisphe- noid rostrum; can cq, cranioquadrate canal; caV, caudal vertebra; ch, secondary choanae; ceri, cervical rib; con spl, contact area of dentary to splenial; con san, contact area from angular to surangular; cond occ, occipital condyle; cpsoc, contact between parietal and supraoccipital; cr, crest; d, dentary; d sel, dorsum sellae; dentt, dentary teeth; dep palp, depression for palpebral attachment; dori, dorsal rib; doV, dorsal vertebra; ec, ectopterygoid; emf, external mandibular PLOS ONE | DOI:10.1371/journal.pone.0160617 February 15, 2017 2 / 47 New atoposaurid crocodyliform from the Upper Jurassic of northwestern Germany fenestra; exoc, exoccipital; f, frontal; fi, fibula; f mag, foramen magnum; fme, external mandib- ular fenestra; for, foramen; for car an, foramen caroticum anterior; for car post, foramen caro- ticum posterior; for eu lat, lateral Eustachian foramen; for eu med, medial Eustachian foramen; for nut, nutricious foramen; for ot, otic foramen; for vag, foramen vagi; for X, fora- men for cranial nerve V; for VI, foramen for cranial nerve VI; for IX-XII, foramina for cranial nerves IX, X, XI, XII; fos cer, cerebral fossa; fos hyp, hypophyseal fossa; hu, humerus; il, ilium; ips, intraparietal suture; itf, infratemporal fenestra; j, jugal; lac, lacrimal; lsph, laterosphenoid; mand, mandible; max, maxilla; mc, metacarpal; mt, metatarsal; n, nasal; na, neural arch; ne, external naris; or, orbit; osd, osteoderm; p, parietal; pal, palatine; palp, palpebral; phal, phalanx; pmx, premaxilla; po, postorbital; prf, prefrontal; prfp, prefrontal pillar; proot, prooticum; pt, pterygoid; pt fl, pterygoid flange; qj, quadratojugal; qu, quadrate; qu cond, quadrate condyle; qu cr A, quadrate crest A; qu cr B, quadrate crest B; rad, radius; san, surangular; scap, scapula; sept, septum; soc, supraoccipital; sof, suborbital fenestra; sp, splenial; sq, squamosal; stf, supra- temporal foramen; symph, mandibular symphysis; ti, tibia; tm1, tooth morph 1; tm2, tooth morph 2. Locality All described material has been found in the Langenberg Quarry at the northern rim of the Harz Mountains near the town of Goslar, Lower Saxony, northern Germany (Fig 1). The quarry exposes well-preserved sections of Upper Jurassic shallow marine strata [29±31]. The beds consist of impure carbonates which grade into marls and which are tilted
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